%PDF-1.3 1 0 obj << /Type /Catalog /Outlines 2 0 R /Pages 3 0 R >> endobj 2 0 obj << /Type /Outlines /Count 0 >> endobj 3 0 obj << /Type /Pages /Kids [6 0 R 82 0 R 164 0 R 198 0 R ] /Count 4 /Resources << /ProcSet 4 0 R /Font << /F1 8 0 R /F2 9 0 R /F3 10 0 R /F4 11 0 R >> /XObject << /I1 12 0 R /I2 13 0 R /I3 34 0 R /I4 35 0 R /I5 56 0 R /I6 57 0 R /I7 78 0 R /I8 79 0 R /I9 80 0 R /I10 81 0 R /I11 104 0 R /I12 105 0 R /I13 130 0 R /I14 131 0 R /I15 156 0 R /I16 157 0 R /I17 162 0 R /I18 163 0 R /I19 168 0 R /I20 169 0 R /I21 178 0 R /I22 179 0 R /I23 188 0 R /I24 189 0 R /I25 196 0 R /I26 197 0 R >> >> /MediaBox [0.000 0.000 612.000 792.000] >> endobj 4 0 obj [/PDF /Text /ImageC ] endobj 5 0 obj << /Creator (DOMPDF) /CreationDate (D:20180722220600+00'00') /ModDate (D:20180722220600+00'00') /Title (Transmission of influenza virus in temperate zones is predominantly by aerosol, in the tropics by contact PLOS Currents Influenza) >> endobj 6 0 obj << /Type /Page /Parent 3 0 R /Annots [ 14 0 R 16 0 R 18 0 R 20 0 R 22 0 R 24 0 R 26 0 R 28 0 R 30 0 R 32 0 R 36 0 R 38 0 R 40 0 R 42 0 R 44 0 R 46 0 R 48 0 R 50 0 R 52 0 R 54 0 R 58 0 R 60 0 R 62 0 R 64 0 R 66 0 R 68 0 R 70 0 R 72 0 R 74 0 R 76 0 R ] /Contents 7 0 R >> endobj 7 0 obj << /Length 19560 >> stream q 375.000 0 0 39.000 222.000 738.000 cm /I2 Do Q q 15.000 662.316 577.500 75.684 re W n 0.267 0.267 0.267 rg BT 15.000 718.042 Td /F2 21.0 Tf [(Transmission of influenza virus in temperate zones is )] TJ ET BT 15.000 693.094 Td /F2 21.0 Tf [(predominantly by aerosol, in the tropics by contact)] TJ ET 0.271 0.267 0.267 rg BT 15.000 671.524 Td /F2 13.5 Tf [(A hypothesis)] TJ ET Q BT 15.000 653.050 Td /F3 9.8 Tf [(August 17, 2009)] TJ ET BT 83.523 653.050 Td /F3 9.8 Tf [()] TJ ET 0.267 0.267 0.267 rg BT 88.398 653.050 Td /F3 9.8 Tf [(Epidemiology)] TJ ET BT 26.250 641.209 Td /F1 9.8 Tf [(Anice Lowen)] TJ ET 0.271 0.267 0.267 rg BT 82.069 641.209 Td /F1 9.8 Tf [(, )] TJ ET 0.267 0.267 0.267 rg BT 87.490 641.209 Td /F1 9.8 Tf [(Peter Palese)] TJ ET 0.271 0.267 0.267 rg BT 26.250 629.304 Td /F1 9.8 Tf [(Lowen A, Palese P. Transmission of influenza virus in temperate zones is predominantly by aerosol, in the tropics by contact: A )] TJ ET BT 26.250 617.400 Td /F1 9.8 Tf [(hypothesis. PLOS Currents Influenza. 2009 Aug 17 . Edition 1. doi: 10.1371/currents.RRN1002.)] TJ ET q 15.000 10.384 577.500 604.634 re W n 0.271 0.267 0.267 rg BT 26.250 588.297 Td /F4 12.0 Tf [(Abstract)] TJ ET BT 26.250 568.343 Td /F1 9.8 Tf [(Using the guinea pig model, we have previously shown that the aerosol transmission of a seasonal human influenza virus is )] TJ ET BT 26.250 556.438 Td /F1 9.8 Tf [(blocked by humid \(80% relative humidity\) or warm \(30C\) ambient conditions. In contrast, we found that transmission by a )] TJ ET BT 26.250 544.533 Td /F1 9.8 Tf [(contact route proceeded at high efficiency despite increased temperature or humidity. Based on these findings, and the )] TJ ET BT 26.250 532.629 Td /F1 9.8 Tf [(observed seasonal behavior of influenza viruses in various regions of the world, we hypothesize herein that the predominant )] TJ ET BT 26.250 520.724 Td /F1 9.8 Tf [(mode of influenza virus transmission differs in temperate and tropical climates. Specifically, we predict that aerosol transmission )] TJ ET BT 26.250 508.819 Td /F1 9.8 Tf [(predominates during the winter season in temperate regions, while contact is the major mode of spread in the tropics. With this )] TJ ET BT 26.250 496.914 Td /F1 9.8 Tf [(idea in mind, possible explanations for the current summer-time spread of swine-origin influenza viruses are discussed.)] TJ ET BT 26.250 460.312 Td /F4 12.0 Tf [(Funding Statement)] TJ ET BT 26.250 440.358 Td /F1 9.8 Tf [(Funding for this work was provided by the Center for Research on Influenza Pathogenesis \(NIAID contract )] TJ ET BT 26.250 428.453 Td /F1 9.8 Tf [(HHSN266200700010C\) and the W. M. Keck Foundation \(to P.P.\). A.C.L. is a Parker B. Francis Fellow in Pulmonary Research.)] TJ ET BT 26.250 399.350 Td /F4 12.0 Tf [(Modes of influenza virus transmission)] TJ ET BT 26.250 379.396 Td /F1 9.8 Tf [(Despite its fundamental importance to controlling the spread of infection, the mode with which influenza viruses transmit from )] TJ ET BT 26.250 367.491 Td /F1 9.8 Tf [(host-to-host remains a point of significant controversy. Three vehicles of viral spread have been implicated: fomites, large )] TJ ET BT 26.250 355.587 Td /F1 9.8 Tf [(respiratory droplets \(usually said to be >10 ?M in diameter\), and small airborne droplet nuclei \(usually said to be < 5 ?M in )] TJ ET BT 26.250 343.682 Td /F1 9.8 Tf [(diameter\) )] TJ ET 0.267 0.267 0.267 rg BT 70.135 345.189 Td /F4 8.7 Tf [(1)] TJ ET 0.271 0.267 0.267 rg BT 74.953 343.682 Td /F1 9.8 Tf [(. Evidence supporting a role for each mode in the transmission of influenza viruses between humans has been )] TJ ET BT 26.250 331.777 Td /F1 9.8 Tf [(reported [reviewed in )] TJ ET 0.267 0.267 0.267 rg BT 119.996 333.284 Td /F4 8.7 Tf [(1)] TJ ET BT 124.815 333.284 Td /F4 8.7 Tf [(2)] TJ ET 0.271 0.267 0.267 rg BT 129.634 331.777 Td /F1 9.8 Tf [(] and under experimental conditions using the guinea pig model all three modes have been observed, )] TJ ET BT 26.250 319.872 Td /F1 9.8 Tf [(albeit with differing efficiencies)] TJ ET 0.267 0.267 0.267 rg BT 157.924 321.380 Td /F4 8.7 Tf [(3)] TJ ET 0.271 0.267 0.267 rg BT 162.742 319.872 Td /F1 9.8 Tf [(. Based on the literature and our own findings, we propose that the possible modes of influenza )] TJ ET BT 26.250 307.968 Td /F1 9.8 Tf [(virus transmission encompass the entire spectrum, from direct contact through to long-range airborne transmission. Which )] TJ ET BT 26.250 296.063 Td /F1 9.8 Tf [(mode of transmission is the most important, or occurs with the highest frequency, is subject to the prevailing circumstances. )] TJ ET BT 26.250 284.158 Td /F1 9.8 Tf [(These circumstances would include the types of surfaces available for fomite transmission, the rate and direction of air )] TJ ET BT 26.250 272.253 Td /F1 9.8 Tf [(exchange, amount of virus shed from the infectious individual, the strain of influenza virus and importantly environmental )] TJ ET BT 26.250 260.349 Td /F1 9.8 Tf [(conditions. The critical question then becomes, not what is the predominant mode of influenza virus transmission, but rather )] TJ ET BT 26.250 248.444 Td /F1 9.8 Tf [(what conditions are favorable to each type of spread? The answer to this question would allow investigation of how variation in )] TJ ET BT 26.250 236.539 Td /F1 9.8 Tf [(mode of transmission with prevailing conditions impacts the epidemiology of influenza. One aspect of this bigger picture in )] TJ ET BT 26.250 224.634 Td /F1 9.8 Tf [(which we are particularly interested is the interplay between the mode of transmission and the seasonality of influenza.)] TJ ET BT 26.250 205.230 Td /F4 9.8 Tf [(Influenza seasonality)] TJ ET BT 26.250 185.825 Td /F1 9.8 Tf [(In temperate regions of the world influenza epidemics occur with a marked winter-time seasonality. Thus, in the Northern )] TJ ET BT 26.250 173.920 Td /F1 9.8 Tf [(Hemisphere, peak influenza activity is seen between November and March, while in the Southern Hemisphere, cases of )] TJ ET BT 26.250 162.015 Td /F1 9.8 Tf [(influenza are most frequent from May to September \( )] TJ ET BT 257.637 162.015 Td /F4 9.8 Tf [(Fig. 1)] TJ ET BT 283.104 162.015 Td /F1 9.8 Tf [( \).)] TJ ET 0.965 0.965 0.965 rg 26.250 10.385 555.000 141.750 re f 0.267 0.267 0.267 rg 0.267 0.267 0.267 RG 26.250 152.135 m 581.250 152.135 l 581.250 151.385 l 26.250 151.385 l f q 225.000 0 0 126.000 35.250 16.385 cm /I4 Do Q q 35.250 10.385 537.000 0.000 re W n Q Q q 15.000 662.316 577.500 75.684 re W n 0.267 0.267 0.267 rg BT 15.000 718.042 Td /F2 21.0 Tf [(Transmission of influenza virus in temperate zones is )] TJ ET BT 15.000 693.094 Td /F2 21.0 Tf [(predominantly by aerosol, in the tropics by contact)] TJ ET 0.271 0.267 0.267 rg BT 15.000 671.524 Td /F2 13.5 Tf [(A hypothesis)] TJ ET Q BT 15.000 653.050 Td /F3 9.8 Tf [(August 17, 2009)] TJ ET BT 83.523 653.050 Td /F3 9.8 Tf [()] TJ ET 0.267 0.267 0.267 rg BT 88.398 653.050 Td /F3 9.8 Tf [(Epidemiology)] TJ ET BT 26.250 641.209 Td /F1 9.8 Tf [(Anice Lowen)] TJ ET 0.271 0.267 0.267 rg BT 82.069 641.209 Td /F1 9.8 Tf [(, )] TJ ET 0.267 0.267 0.267 rg BT 87.490 641.209 Td /F1 9.8 Tf [(Peter Palese)] TJ ET 0.271 0.267 0.267 rg BT 26.250 629.304 Td /F1 9.8 Tf [(Lowen A, Palese P. Transmission of influenza virus in temperate zones is predominantly by aerosol, in the tropics by contact: A )] TJ ET BT 26.250 617.400 Td /F1 9.8 Tf [(hypothesis. PLOS Currents Influenza. 2009 Aug 17 . Edition 1. doi: 10.1371/currents.RRN1002.)] TJ ET q 15.000 10.384 577.500 604.634 re W n 0.271 0.267 0.267 rg BT 26.250 588.297 Td /F4 12.0 Tf [(Abstract)] TJ ET BT 26.250 568.343 Td /F1 9.8 Tf [(Using the guinea pig model, we have previously shown that the aerosol transmission of a seasonal human influenza virus is )] TJ ET BT 26.250 556.438 Td /F1 9.8 Tf [(blocked by humid \(80% relative humidity\) or warm \(30C\) ambient conditions. In contrast, we found that transmission by a )] TJ ET BT 26.250 544.533 Td /F1 9.8 Tf [(contact route proceeded at high efficiency despite increased temperature or humidity. Based on these findings, and the )] TJ ET BT 26.250 532.629 Td /F1 9.8 Tf [(observed seasonal behavior of influenza viruses in various regions of the world, we hypothesize herein that the predominant )] TJ ET BT 26.250 520.724 Td /F1 9.8 Tf [(mode of influenza virus transmission differs in temperate and tropical climates. Specifically, we predict that aerosol transmission )] TJ ET BT 26.250 508.819 Td /F1 9.8 Tf [(predominates during the winter season in temperate regions, while contact is the major mode of spread in the tropics. With this )] TJ ET BT 26.250 496.914 Td /F1 9.8 Tf [(idea in mind, possible explanations for the current summer-time spread of swine-origin influenza viruses are discussed.)] TJ ET BT 26.250 460.312 Td /F4 12.0 Tf [(Funding Statement)] TJ ET BT 26.250 440.358 Td /F1 9.8 Tf [(Funding for this work was provided by the Center for Research on Influenza Pathogenesis \(NIAID contract )] TJ ET BT 26.250 428.453 Td /F1 9.8 Tf [(HHSN266200700010C\) and the W. M. Keck Foundation \(to P.P.\). A.C.L. is a Parker B. Francis Fellow in Pulmonary Research.)] TJ ET BT 26.250 399.350 Td /F4 12.0 Tf [(Modes of influenza virus transmission)] TJ ET BT 26.250 379.396 Td /F1 9.8 Tf [(Despite its fundamental importance to controlling the spread of infection, the mode with which influenza viruses transmit from )] TJ ET BT 26.250 367.491 Td /F1 9.8 Tf [(host-to-host remains a point of significant controversy. Three vehicles of viral spread have been implicated: fomites, large )] TJ ET BT 26.250 355.587 Td /F1 9.8 Tf [(respiratory droplets \(usually said to be >10 ?M in diameter\), and small airborne droplet nuclei \(usually said to be < 5 ?M in )] TJ ET BT 26.250 343.682 Td /F1 9.8 Tf [(diameter\) )] TJ ET 0.267 0.267 0.267 rg BT 70.135 345.189 Td /F4 8.7 Tf [(1)] TJ ET 0.271 0.267 0.267 rg BT 74.953 343.682 Td /F1 9.8 Tf [(. Evidence supporting a role for each mode in the transmission of influenza viruses between humans has been )] TJ ET BT 26.250 331.777 Td /F1 9.8 Tf [(reported [reviewed in )] TJ ET 0.267 0.267 0.267 rg BT 119.996 333.284 Td /F4 8.7 Tf [(1)] TJ ET BT 124.815 333.284 Td /F4 8.7 Tf [(2)] TJ ET 0.271 0.267 0.267 rg BT 129.634 331.777 Td /F1 9.8 Tf [(] and under experimental conditions using the guinea pig model all three modes have been observed, )] TJ ET BT 26.250 319.872 Td /F1 9.8 Tf [(albeit with differing efficiencies)] TJ ET 0.267 0.267 0.267 rg BT 157.924 321.380 Td /F4 8.7 Tf [(3)] TJ ET 0.271 0.267 0.267 rg BT 162.742 319.872 Td /F1 9.8 Tf [(. Based on the literature and our own findings, we propose that the possible modes of influenza )] TJ ET BT 26.250 307.968 Td /F1 9.8 Tf [(virus transmission encompass the entire spectrum, from direct contact through to long-range airborne transmission. Which )] TJ ET BT 26.250 296.063 Td /F1 9.8 Tf [(mode of transmission is the most important, or occurs with the highest frequency, is subject to the prevailing circumstances. )] TJ ET BT 26.250 284.158 Td /F1 9.8 Tf [(These circumstances would include the types of surfaces available for fomite transmission, the rate and direction of air )] TJ ET BT 26.250 272.253 Td /F1 9.8 Tf [(exchange, amount of virus shed from the infectious individual, the strain of influenza virus and importantly environmental )] TJ ET BT 26.250 260.349 Td /F1 9.8 Tf [(conditions. The critical question then becomes, not what is the predominant mode of influenza virus transmission, but rather )] TJ ET BT 26.250 248.444 Td /F1 9.8 Tf [(what conditions are favorable to each type of spread? The answer to this question would allow investigation of how variation in )] TJ ET BT 26.250 236.539 Td /F1 9.8 Tf [(mode of transmission with prevailing conditions impacts the epidemiology of influenza. One aspect of this bigger picture in )] TJ ET BT 26.250 224.634 Td /F1 9.8 Tf [(which we are particularly interested is the interplay between the mode of transmission and the seasonality of influenza.)] TJ ET BT 26.250 205.230 Td /F4 9.8 Tf [(Influenza seasonality)] TJ ET BT 26.250 185.825 Td /F1 9.8 Tf [(In temperate regions of the world influenza epidemics occur with a marked winter-time seasonality. Thus, in the Northern )] TJ ET BT 26.250 173.920 Td /F1 9.8 Tf [(Hemisphere, peak influenza activity is seen between November and March, while in the Southern Hemisphere, cases of )] TJ ET BT 26.250 162.015 Td /F1 9.8 Tf [(influenza are most frequent from May to September \( )] TJ ET BT 257.637 162.015 Td /F4 9.8 Tf [(Fig. 1)] TJ ET BT 283.104 162.015 Td /F1 9.8 Tf [( \).)] TJ ET 0.965 0.965 0.965 rg 26.250 10.385 555.000 141.750 re f 0.267 0.267 0.267 rg 0.267 0.267 0.267 RG 26.250 152.135 m 581.250 152.135 l 581.250 151.385 l 26.250 151.385 l f q 225.000 0 0 126.000 35.250 16.385 cm /I6 Do Q q 35.250 10.385 537.000 0.000 re W n Q Q q 15.000 662.316 577.500 75.684 re W n 0.267 0.267 0.267 rg BT 15.000 718.042 Td /F2 21.0 Tf [(Transmission of influenza virus in temperate zones is )] TJ ET BT 15.000 693.094 Td /F2 21.0 Tf [(predominantly by aerosol, in the tropics by contact)] TJ ET 0.271 0.267 0.267 rg BT 15.000 671.524 Td /F2 13.5 Tf [(A hypothesis)] TJ ET Q BT 15.000 653.050 Td /F3 9.8 Tf [(August 17, 2009)] TJ ET BT 83.523 653.050 Td /F3 9.8 Tf [()] TJ ET 0.267 0.267 0.267 rg BT 88.398 653.050 Td /F3 9.8 Tf [(Epidemiology)] TJ ET BT 26.250 641.209 Td /F1 9.8 Tf [(Anice Lowen)] TJ ET 0.271 0.267 0.267 rg BT 82.069 641.209 Td /F1 9.8 Tf [(, )] TJ ET 0.267 0.267 0.267 rg BT 87.490 641.209 Td /F1 9.8 Tf [(Peter Palese)] TJ ET 0.271 0.267 0.267 rg BT 26.250 629.304 Td /F1 9.8 Tf [(Lowen A, Palese P. Transmission of influenza virus in temperate zones is predominantly by aerosol, in the tropics by contact: A )] TJ ET BT 26.250 617.400 Td /F1 9.8 Tf [(hypothesis. PLOS Currents Influenza. 2009 Aug 17 . Edition 1. doi: 10.1371/currents.RRN1002.)] TJ ET q 15.000 10.384 577.500 604.634 re W n 0.271 0.267 0.267 rg BT 26.250 588.297 Td /F4 12.0 Tf [(Abstract)] TJ ET BT 26.250 568.343 Td /F1 9.8 Tf [(Using the guinea pig model, we have previously shown that the aerosol transmission of a seasonal human influenza virus is )] TJ ET BT 26.250 556.438 Td /F1 9.8 Tf [(blocked by humid \(80% relative humidity\) or warm \(30C\) ambient conditions. In contrast, we found that transmission by a )] TJ ET BT 26.250 544.533 Td /F1 9.8 Tf [(contact route proceeded at high efficiency despite increased temperature or humidity. Based on these findings, and the )] TJ ET BT 26.250 532.629 Td /F1 9.8 Tf [(observed seasonal behavior of influenza viruses in various regions of the world, we hypothesize herein that the predominant )] TJ ET BT 26.250 520.724 Td /F1 9.8 Tf [(mode of influenza virus transmission differs in temperate and tropical climates. Specifically, we predict that aerosol transmission )] TJ ET BT 26.250 508.819 Td /F1 9.8 Tf [(predominates during the winter season in temperate regions, while contact is the major mode of spread in the tropics. With this )] TJ ET BT 26.250 496.914 Td /F1 9.8 Tf [(idea in mind, possible explanations for the current summer-time spread of swine-origin influenza viruses are discussed.)] TJ ET BT 26.250 460.312 Td /F4 12.0 Tf [(Funding Statement)] TJ ET BT 26.250 440.358 Td /F1 9.8 Tf [(Funding for this work was provided by the Center for Research on Influenza Pathogenesis \(NIAID contract )] TJ ET BT 26.250 428.453 Td /F1 9.8 Tf [(HHSN266200700010C\) and the W. M. Keck Foundation \(to P.P.\). A.C.L. is a Parker B. Francis Fellow in Pulmonary Research.)] TJ ET BT 26.250 399.350 Td /F4 12.0 Tf [(Modes of influenza virus transmission)] TJ ET BT 26.250 379.396 Td /F1 9.8 Tf [(Despite its fundamental importance to controlling the spread of infection, the mode with which influenza viruses transmit from )] TJ ET BT 26.250 367.491 Td /F1 9.8 Tf [(host-to-host remains a point of significant controversy. Three vehicles of viral spread have been implicated: fomites, large )] TJ ET BT 26.250 355.587 Td /F1 9.8 Tf [(respiratory droplets \(usually said to be >10 ?M in diameter\), and small airborne droplet nuclei \(usually said to be < 5 ?M in )] TJ ET BT 26.250 343.682 Td /F1 9.8 Tf [(diameter\) )] TJ ET 0.267 0.267 0.267 rg BT 70.135 345.189 Td /F4 8.7 Tf [(1)] TJ ET 0.271 0.267 0.267 rg BT 74.953 343.682 Td /F1 9.8 Tf [(. Evidence supporting a role for each mode in the transmission of influenza viruses between humans has been )] TJ ET BT 26.250 331.777 Td /F1 9.8 Tf [(reported [reviewed in )] TJ ET 0.267 0.267 0.267 rg BT 119.996 333.284 Td /F4 8.7 Tf [(1)] TJ ET BT 124.815 333.284 Td /F4 8.7 Tf [(2)] TJ ET 0.271 0.267 0.267 rg BT 129.634 331.777 Td /F1 9.8 Tf [(] and under experimental conditions using the guinea pig model all three modes have been observed, )] TJ ET BT 26.250 319.872 Td /F1 9.8 Tf [(albeit with differing efficiencies)] TJ ET 0.267 0.267 0.267 rg BT 157.924 321.380 Td /F4 8.7 Tf [(3)] TJ ET 0.271 0.267 0.267 rg BT 162.742 319.872 Td /F1 9.8 Tf [(. Based on the literature and our own findings, we propose that the possible modes of influenza )] TJ ET BT 26.250 307.968 Td /F1 9.8 Tf [(virus transmission encompass the entire spectrum, from direct contact through to long-range airborne transmission. Which )] TJ ET BT 26.250 296.063 Td /F1 9.8 Tf [(mode of transmission is the most important, or occurs with the highest frequency, is subject to the prevailing circumstances. )] TJ ET BT 26.250 284.158 Td /F1 9.8 Tf [(These circumstances would include the types of surfaces available for fomite transmission, the rate and direction of air )] TJ ET BT 26.250 272.253 Td /F1 9.8 Tf [(exchange, amount of virus shed from the infectious individual, the strain of influenza virus and importantly environmental )] TJ ET BT 26.250 260.349 Td /F1 9.8 Tf [(conditions. The critical question then becomes, not what is the predominant mode of influenza virus transmission, but rather )] TJ ET BT 26.250 248.444 Td /F1 9.8 Tf [(what conditions are favorable to each type of spread? The answer to this question would allow investigation of how variation in )] TJ ET BT 26.250 236.539 Td /F1 9.8 Tf [(mode of transmission with prevailing conditions impacts the epidemiology of influenza. One aspect of this bigger picture in )] TJ ET BT 26.250 224.634 Td /F1 9.8 Tf [(which we are particularly interested is the interplay between the mode of transmission and the seasonality of influenza.)] TJ ET BT 26.250 205.230 Td /F4 9.8 Tf [(Influenza seasonality)] TJ ET BT 26.250 185.825 Td /F1 9.8 Tf [(In temperate regions of the world influenza epidemics occur with a marked winter-time seasonality. Thus, in the Northern )] TJ ET BT 26.250 173.920 Td /F1 9.8 Tf [(Hemisphere, peak influenza activity is seen between November and March, while in the Southern Hemisphere, cases of )] TJ ET BT 26.250 162.015 Td /F1 9.8 Tf [(influenza are most frequent from May to September \( )] TJ ET BT 257.637 162.015 Td /F4 9.8 Tf [(Fig. 1)] TJ ET BT 283.104 162.015 Td /F1 9.8 Tf [( \).)] TJ ET 0.965 0.965 0.965 rg 26.250 10.385 555.000 141.750 re f 0.267 0.267 0.267 rg 0.267 0.267 0.267 RG 26.250 152.135 m 581.250 152.135 l 581.250 151.385 l 26.250 151.385 l f q 225.000 0 0 126.000 35.250 16.385 cm /I8 Do Q q 35.250 10.385 537.000 0.000 re W n Q Q q 225.000 0 0 126.000 35.250 16.385 cm /I10 Do Q q 0.000 0.000 0.000 rg BT 291.710 19.825 Td /F1 11.0 Tf [(1)] TJ ET BT 25.000 19.825 Td /F1 11.0 Tf [(PLOS Currents Influenza)] TJ ET Q endstream endobj 8 0 obj << /Type /Font /Subtype /Type1 /Name /F1 /BaseFont /Helvetica /Encoding /WinAnsiEncoding >> endobj 9 0 obj << /Type /Font /Subtype /Type1 /Name /F2 /BaseFont /Times-Bold /Encoding /WinAnsiEncoding >> endobj 10 0 obj << /Type /Font /Subtype /Type1 /Name /F3 /BaseFont /Times-Italic /Encoding /WinAnsiEncoding >> endobj 11 0 obj << /Type /Font /Subtype /Type1 /Name /F4 /BaseFont /Helvetica-Bold /Encoding /WinAnsiEncoding >> endobj 12 0 obj << /Type /XObject /Subtype /Image /Width 500 /Height 52 /Filter /FlateDecode /DecodeParms << /Predictor 15 /Colors 1 /Columns 500 /BitsPerComponent 8>> /ColorSpace /DeviceGray /BitsPerComponent 8 /Length 144>> stream x1 0 'ݲ؎"e{dzAdzAdzAdzAdzAdzAdzAdzAdzAdzAdzAdzAtlM0\ endstream endobj 13 0 obj << /Type /XObject /Subtype /Image /Width 500 /Height 52 /SMask 12 0 R /Filter /FlateDecode /DecodeParms << /Predictor 15 /Colors 3 /Columns 500 /BitsPerComponent 8>> /ColorSpace /DeviceRGB /BitsPerComponent 8 /Length 3574>> stream x}ƕK*8+0T ֮ S L*0߇d" sl2FBiFwY0 00 0ŝa0 aqgR܋x||lu2 03UL۶BxI 0:UU0x;֝($z%șHڶ ÐVz ]IS> â(.* TQUuL5 $>BvpH0U:xkkGJT{OɗRNyGOΙyn)q7`(~_Ji2vgk]qVꔷ3N'ȴy4Ikm6!q:FJ)-0n܋v!^}_Z l61va_~Bxw8mv;EJY8}zzQ3]Pȹ,2_jfY4MQ[ '.uO O͋kշ9b6.#ߦUUzhf09+nKɿ4 f7Bu7ĨMfrF0`s8429j1}ZWDW;A4ҫ'TQ{7Cb[$ -"ΨMQgzN/2R7Tco*W:N݋6TUu8`RSi O>r;z^m(0w`:m=]գCE&t |ZYQ%{X[nR+zbEFΧ5pzq`ުం Ih{9G 4wMx' nmvEZ sg7?ZQ}&ҎKGuPu.j8UgNKꓙQC)s#$1䧈x[Z(|Xzc;t<~50Hh_T3Ur>a$]B8Fz[?K>Y/rQ=E4?EaLqzpX!(nNut]}e(/S\fz,<ZOEa]0 sCS_}I Y}n۶r(fg7#)h6ա}$ üvq]Aא$c0ddBDQ)ueypU-z~9Hk<$I©y'Tk`yY d}e{䨙ǐu>&w񚓶('GI\rm[G4 Sk7$B pEkYYC/ 9&Ʋ[V9MQ':X|C֨ʉ jt"ޡ+rttBռGdiM FuTEi(łv]fk"8Ldp' ^הMup2`M?dE!|KJ-\.MzکC=&¾HfR7$ev,La0+$OFuEQ"i,ːN?~E*t4qΚF:t`,{RkR]ޜ(Ԕѣ$IbN}5لS1(Ȱ$I$(eWp8Lٱ3߯ky۶&. 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X>[,Yƹ-CSe:5_hc#T(&Z hT?dž/(<-gfH:oS1wlݶi6+^W=Uk_@ &p7an(#v)b aRaK.|?D+-Y{JC0BPᱟoo,FK.QʒIYg#g7> endobj 83 0 obj << /Length 22065 >> stream q 15.000 48.350 577.500 728.650 re W n 0.965 0.965 0.965 rg 26.250 699.136 555.000 77.864 re f 0.267 0.267 0.267 rg 0.267 0.267 0.267 RG 26.250 699.136 m 581.250 699.136 l 581.250 699.886 l 26.250 699.886 l f q 35.250 710.386 537.000 66.614 re W n 0.271 0.267 0.267 rg BT 35.250 767.476 Td /F4 9.8 Tf [(Fig. 1: Weekly influenza activity for five countries spanning the Americas.)] TJ ET BT 35.250 748.106 Td /F1 9.8 Tf [(Adapted from reference )] TJ ET 0.267 0.267 0.267 rg BT 140.384 751.079 Td /F4 8.7 Tf [(4)] TJ ET 0.271 0.267 0.267 rg BT 145.203 748.106 Td /F1 9.8 Tf [( , this graph illustrates the differing patterns of influenza activity seen in temperate and tropical )] TJ ET BT 35.250 734.370 Td /F1 9.8 Tf [(\(here exemplified by Colombia\) regions. The latitude of the capital city of each country is indicated in brackets in the figure )] TJ ET BT 35.250 720.634 Td /F1 9.8 Tf [(legend.)] TJ ET Q BT 26.250 682.113 Td /F1 9.8 Tf [(In the tropics, by contrast, influenza activity tends to be much more sporadic, with isolations occurring either throughout the year )] TJ ET BT 26.250 670.208 Td /F1 9.8 Tf [(or in temporally irregular outbreaks \(Fig. 1\) )] TJ ET 0.267 0.267 0.267 rg BT 213.177 671.715 Td /F4 8.7 Tf [(4)] TJ ET 0.271 0.267 0.267 rg BT 217.996 670.208 Td /F1 9.8 Tf [( . For many years the underlying reasons for influenza seasonality were widely )] TJ ET BT 26.250 658.303 Td /F1 9.8 Tf [(discussed but poorly understood )] TJ ET 0.267 0.267 0.267 rg BT 169.321 659.810 Td /F4 8.7 Tf [(5)] TJ ET BT 174.140 659.810 Td /F4 8.7 Tf [(6)] TJ ET 0.271 0.267 0.267 rg BT 178.959 658.303 Td /F1 9.8 Tf [(; recently, however, we have reported compelling evidence obtained in the guinea pig model )] TJ ET BT 26.250 646.398 Td /F1 9.8 Tf [(which suggests that seasonal variation in humidity and temperature may direct seasonal fluctuations in influenza activity )] TJ ET 0.267 0.267 0.267 rg BT 545.389 647.906 Td /F4 8.7 Tf [(7)] TJ ET BT 550.207 647.906 Td /F4 8.7 Tf [(8)] TJ ET 0.271 0.267 0.267 rg BT 555.026 646.398 Td /F1 9.8 Tf [( . In )] TJ ET BT 26.250 634.494 Td /F1 9.8 Tf [(addition, as demonstrated below, our data revealed that transmission by the aerosol route \(here defined as encompassing both )] TJ ET BT 26.250 622.589 Td /F1 9.8 Tf [(large and small droplet spread\) is acutely sensitive to both humidity and temperature, while transmission by the contact route is )] TJ ET BT 26.250 610.684 Td /F1 9.8 Tf [(not.)] TJ ET BT 26.250 591.279 Td /F4 9.8 Tf [(Sensitivity of aerosol transmission to humidity and temperature)] TJ ET BT 26.250 571.875 Td /F1 9.8 Tf [(By combining the guinea pig transmission model with the use of an environmentally controlled chamber, we have been able to )] TJ ET BT 26.250 559.970 Td /F1 9.8 Tf [(systematically evaluate the effects of temperature and relative humidity on influenza virus transmission )] TJ ET 0.267 0.267 0.267 rg BT 471.133 561.477 Td /F4 8.7 Tf [(7)] TJ ET BT 475.951 561.477 Td /F4 8.7 Tf [(8)] TJ ET 0.271 0.267 0.267 rg BT 480.770 559.970 Td /F1 9.8 Tf [( . To specifically test )] TJ ET BT 26.250 548.065 Td /F1 9.8 Tf [(the effects on transmission by the aerosol route, we used an experimental design in which infected and exposed guinea pigs )] TJ ET BT 26.250 536.160 Td /F1 9.8 Tf [(were placed in separate cages )] TJ ET 0.267 0.267 0.267 rg BT 161.190 537.668 Td /F4 8.7 Tf [(7)] TJ ET 0.271 0.267 0.267 rg BT 166.009 536.160 Td /F1 9.8 Tf [(. Each cage was open to air flow on the top and on one side and pairs of cages were placed at )] TJ ET BT 26.250 524.256 Td /F1 9.8 Tf [(a minimum distance of approximately 2 cm such that the guinea pigs were not able to touch each other. Following this )] TJ ET BT 26.250 512.351 Td /F1 9.8 Tf [(arrangement, four infected and four nave animals were placed together into an environmental chamber and monitored for virus )] TJ ET BT 26.250 500.446 Td /F1 9.8 Tf [(shedding over a period of one week. This strategy revealed a strong impact of ambient temperature and humidity conditions on )] TJ ET BT 26.250 488.541 Td /F1 9.8 Tf [(the aerosol transmission of influenza viruses \( )] TJ ET BT 226.739 488.541 Td /F4 9.8 Tf [(Fig. 2)] TJ ET BT 252.206 488.541 Td /F1 9.8 Tf [( \). In general, we found that cold and dry conditions were most favorable, )] TJ ET BT 26.250 476.637 Td /F1 9.8 Tf [(while aerosol transmission was completely blocked by warm \(30C\) or humid \(80% relative humidity\) conditions.)] TJ ET 0.965 0.965 0.965 rg 26.250 154.183 555.000 312.572 re f 0.267 0.267 0.267 rg 0.267 0.267 0.267 RG 26.250 466.756 m 581.250 466.756 l 581.250 466.006 l 26.250 466.006 l f 26.250 154.183 m 581.250 154.183 l 581.250 154.933 l 26.250 154.933 l f q 225.000 0 0 177.750 35.250 279.256 cm /I12 Do Q q 35.250 165.433 537.000 107.822 re W n 0.271 0.267 0.267 rg BT 35.250 263.732 Td /F4 9.8 Tf [(Fig. 2: Aerosol transmission of influenza A/Panama/2007/1999 virus varies with relative humidity and temperature.)] TJ ET BT 35.250 244.362 Td /F1 9.8 Tf [(Nasal wash titers of intranasally inoculated animals are indicated by dashed lines, while nasal wash titers of exposed )] TJ ET BT 35.250 230.626 Td /F1 9.8 Tf [(animals are shown with solid lines. In each experiment, n = 4 inoculated guinea pigs and 4 exposed guinea pigs. The )] TJ ET BT 35.250 216.889 Td /F1 9.8 Tf [(environmental conditions under which each experiment was performed are indicated at the top \(temperature\) and at the left )] TJ ET BT 35.250 203.153 Td /F1 9.8 Tf [(\(relative humidity, RH\). Comparison of results across each row shows the decrease in aerosol transmission with increasing )] TJ ET BT 35.250 189.417 Td /F1 9.8 Tf [(temperature; comparison of results down each column indicates the decrease in aerosol transmission with the increase in )] TJ ET BT 35.250 175.681 Td /F1 9.8 Tf [(relative humidity )] TJ ET 0.267 0.267 0.267 rg BT 108.394 178.653 Td /F4 8.7 Tf [(7)] TJ ET 0.271 0.267 0.267 rg BT 113.213 175.681 Td /F1 9.8 Tf [( .)] TJ ET Q BT 26.250 137.160 Td /F4 9.8 Tf [(Insensitivity of contact transmission to humidity and temperature)] TJ ET BT 26.250 117.755 Td /F1 9.8 Tf [(To evaluate whether transmission by the contact route would also be affected by humidity and temperature, we used an )] TJ ET BT 26.250 105.850 Td /F1 9.8 Tf [(experimental design in which infected and exposed guinea pigs were placed in the same cage together )] TJ ET 0.267 0.267 0.267 rg BT 472.810 107.357 Td /F4 8.7 Tf [(8)] TJ ET 0.271 0.267 0.267 rg BT 477.628 105.850 Td /F1 9.8 Tf [(. This set up allows for )] TJ ET BT 26.250 93.945 Td /F1 9.8 Tf [(transmission to occur by direct or indirect contact, as well as by short-range aerosol transmission. As with the aerosol )] TJ ET BT 26.250 82.041 Td /F1 9.8 Tf [(transmission experiments, environmental conditions were controlled within a chamber and exposure occurred over a period of )] TJ ET BT 26.250 70.136 Td /F1 9.8 Tf [(one week. As shown in )] TJ ET BT 128.674 70.136 Td /F4 9.8 Tf [(Figure 3)] TJ ET BT 166.601 70.136 Td /F1 9.8 Tf [( , these experiments indicated that the efficiency of transmission by the contact route varies )] TJ ET BT 26.250 58.231 Td /F1 9.8 Tf [(very little with relative humidity or temperature.)] TJ ET Q q 15.000 48.350 577.500 728.650 re W n 0.965 0.965 0.965 rg 26.250 699.136 555.000 77.864 re f 0.267 0.267 0.267 rg 0.267 0.267 0.267 RG 26.250 699.136 m 581.250 699.136 l 581.250 699.886 l 26.250 699.886 l f q 35.250 710.386 537.000 66.614 re W n 0.271 0.267 0.267 rg BT 35.250 767.476 Td /F4 9.8 Tf [(Fig. 1: Weekly influenza activity for five countries spanning the Americas.)] TJ ET BT 35.250 748.106 Td /F1 9.8 Tf [(Adapted from reference )] TJ ET 0.267 0.267 0.267 rg BT 140.384 751.079 Td /F4 8.7 Tf [(4)] TJ ET 0.271 0.267 0.267 rg BT 145.203 748.106 Td /F1 9.8 Tf [( , this graph illustrates the differing patterns of influenza activity seen in temperate and tropical )] TJ ET BT 35.250 734.370 Td /F1 9.8 Tf [(\(here exemplified by Colombia\) regions. The latitude of the capital city of each country is indicated in brackets in the figure )] TJ ET BT 35.250 720.634 Td /F1 9.8 Tf [(legend.)] TJ ET Q BT 26.250 682.113 Td /F1 9.8 Tf [(In the tropics, by contrast, influenza activity tends to be much more sporadic, with isolations occurring either throughout the year )] TJ ET BT 26.250 670.208 Td /F1 9.8 Tf [(or in temporally irregular outbreaks \(Fig. 1\) )] TJ ET 0.267 0.267 0.267 rg BT 213.177 671.715 Td /F4 8.7 Tf [(4)] TJ ET 0.271 0.267 0.267 rg BT 217.996 670.208 Td /F1 9.8 Tf [( . For many years the underlying reasons for influenza seasonality were widely )] TJ ET BT 26.250 658.303 Td /F1 9.8 Tf [(discussed but poorly understood )] TJ ET 0.267 0.267 0.267 rg BT 169.321 659.810 Td /F4 8.7 Tf [(5)] TJ ET BT 174.140 659.810 Td /F4 8.7 Tf [(6)] TJ ET 0.271 0.267 0.267 rg BT 178.959 658.303 Td /F1 9.8 Tf [(; recently, however, we have reported compelling evidence obtained in the guinea pig model )] TJ ET BT 26.250 646.398 Td /F1 9.8 Tf [(which suggests that seasonal variation in humidity and temperature may direct seasonal fluctuations in influenza activity )] TJ ET 0.267 0.267 0.267 rg BT 545.389 647.906 Td /F4 8.7 Tf [(7)] TJ ET BT 550.207 647.906 Td /F4 8.7 Tf [(8)] TJ ET 0.271 0.267 0.267 rg BT 555.026 646.398 Td /F1 9.8 Tf [( . In )] TJ ET BT 26.250 634.494 Td /F1 9.8 Tf [(addition, as demonstrated below, our data revealed that transmission by the aerosol route \(here defined as encompassing both )] TJ ET BT 26.250 622.589 Td /F1 9.8 Tf [(large and small droplet spread\) is acutely sensitive to both humidity and temperature, while transmission by the contact route is )] TJ ET BT 26.250 610.684 Td /F1 9.8 Tf [(not.)] TJ ET BT 26.250 591.279 Td /F4 9.8 Tf [(Sensitivity of aerosol transmission to humidity and temperature)] TJ ET BT 26.250 571.875 Td /F1 9.8 Tf [(By combining the guinea pig transmission model with the use of an environmentally controlled chamber, we have been able to )] TJ ET BT 26.250 559.970 Td /F1 9.8 Tf [(systematically evaluate the effects of temperature and relative humidity on influenza virus transmission )] TJ ET 0.267 0.267 0.267 rg BT 471.133 561.477 Td /F4 8.7 Tf [(7)] TJ ET BT 475.951 561.477 Td /F4 8.7 Tf [(8)] TJ ET 0.271 0.267 0.267 rg BT 480.770 559.970 Td /F1 9.8 Tf [( . To specifically test )] TJ ET BT 26.250 548.065 Td /F1 9.8 Tf [(the effects on transmission by the aerosol route, we used an experimental design in which infected and exposed guinea pigs )] TJ ET BT 26.250 536.160 Td /F1 9.8 Tf [(were placed in separate cages )] TJ ET 0.267 0.267 0.267 rg BT 161.190 537.668 Td /F4 8.7 Tf [(7)] TJ ET 0.271 0.267 0.267 rg BT 166.009 536.160 Td /F1 9.8 Tf [(. Each cage was open to air flow on the top and on one side and pairs of cages were placed at )] TJ ET BT 26.250 524.256 Td /F1 9.8 Tf [(a minimum distance of approximately 2 cm such that the guinea pigs were not able to touch each other. Following this )] TJ ET BT 26.250 512.351 Td /F1 9.8 Tf [(arrangement, four infected and four nave animals were placed together into an environmental chamber and monitored for virus )] TJ ET BT 26.250 500.446 Td /F1 9.8 Tf [(shedding over a period of one week. This strategy revealed a strong impact of ambient temperature and humidity conditions on )] TJ ET BT 26.250 488.541 Td /F1 9.8 Tf [(the aerosol transmission of influenza viruses \( )] TJ ET BT 226.739 488.541 Td /F4 9.8 Tf [(Fig. 2)] TJ ET BT 252.206 488.541 Td /F1 9.8 Tf [( \). In general, we found that cold and dry conditions were most favorable, )] TJ ET BT 26.250 476.637 Td /F1 9.8 Tf [(while aerosol transmission was completely blocked by warm \(30C\) or humid \(80% relative humidity\) conditions.)] TJ ET 0.965 0.965 0.965 rg 26.250 154.183 555.000 312.572 re f 0.267 0.267 0.267 rg 0.267 0.267 0.267 RG 26.250 466.756 m 581.250 466.756 l 581.250 466.006 l 26.250 466.006 l f 26.250 154.183 m 581.250 154.183 l 581.250 154.933 l 26.250 154.933 l f q 225.000 0 0 177.750 35.250 279.256 cm /I14 Do Q q 35.250 165.433 537.000 107.822 re W n 0.271 0.267 0.267 rg BT 35.250 263.732 Td /F4 9.8 Tf [(Fig. 2: Aerosol transmission of influenza A/Panama/2007/1999 virus varies with relative humidity and temperature.)] TJ ET BT 35.250 244.362 Td /F1 9.8 Tf [(Nasal wash titers of intranasally inoculated animals are indicated by dashed lines, while nasal wash titers of exposed )] TJ ET BT 35.250 230.626 Td /F1 9.8 Tf [(animals are shown with solid lines. In each experiment, n = 4 inoculated guinea pigs and 4 exposed guinea pigs. The )] TJ ET BT 35.250 216.889 Td /F1 9.8 Tf [(environmental conditions under which each experiment was performed are indicated at the top \(temperature\) and at the left )] TJ ET BT 35.250 203.153 Td /F1 9.8 Tf [(\(relative humidity, RH\). Comparison of results across each row shows the decrease in aerosol transmission with increasing )] TJ ET BT 35.250 189.417 Td /F1 9.8 Tf [(temperature; comparison of results down each column indicates the decrease in aerosol transmission with the increase in )] TJ ET BT 35.250 175.681 Td /F1 9.8 Tf [(relative humidity )] TJ ET 0.267 0.267 0.267 rg BT 108.394 178.653 Td /F4 8.7 Tf [(7)] TJ ET 0.271 0.267 0.267 rg BT 113.213 175.681 Td /F1 9.8 Tf [( .)] TJ ET Q BT 26.250 137.160 Td /F4 9.8 Tf [(Insensitivity of contact transmission to humidity and temperature)] TJ ET BT 26.250 117.755 Td /F1 9.8 Tf [(To evaluate whether transmission by the contact route would also be affected by humidity and temperature, we used an )] TJ ET BT 26.250 105.850 Td /F1 9.8 Tf [(experimental design in which infected and exposed guinea pigs were placed in the same cage together )] TJ ET 0.267 0.267 0.267 rg BT 472.810 107.357 Td /F4 8.7 Tf [(8)] TJ ET 0.271 0.267 0.267 rg BT 477.628 105.850 Td /F1 9.8 Tf [(. This set up allows for )] TJ ET BT 26.250 93.945 Td /F1 9.8 Tf [(transmission to occur by direct or indirect contact, as well as by short-range aerosol transmission. As with the aerosol )] TJ ET BT 26.250 82.041 Td /F1 9.8 Tf [(transmission experiments, environmental conditions were controlled within a chamber and exposure occurred over a period of )] TJ ET BT 26.250 70.136 Td /F1 9.8 Tf [(one week. As shown in )] TJ ET BT 128.674 70.136 Td /F4 9.8 Tf [(Figure 3)] TJ ET BT 166.601 70.136 Td /F1 9.8 Tf [( , these experiments indicated that the efficiency of transmission by the contact route varies )] TJ ET BT 26.250 58.231 Td /F1 9.8 Tf [(very little with relative humidity or temperature.)] TJ ET Q q 15.000 48.350 577.500 728.650 re W n 0.965 0.965 0.965 rg 26.250 699.136 555.000 77.864 re f 0.267 0.267 0.267 rg 0.267 0.267 0.267 RG 26.250 699.136 m 581.250 699.136 l 581.250 699.886 l 26.250 699.886 l f q 35.250 710.386 537.000 66.614 re W n 0.271 0.267 0.267 rg BT 35.250 767.476 Td /F4 9.8 Tf [(Fig. 1: Weekly influenza activity for five countries spanning the Americas.)] TJ ET BT 35.250 748.106 Td /F1 9.8 Tf [(Adapted from reference )] TJ ET 0.267 0.267 0.267 rg BT 140.384 751.079 Td /F4 8.7 Tf [(4)] TJ ET 0.271 0.267 0.267 rg BT 145.203 748.106 Td /F1 9.8 Tf [( , this graph illustrates the differing patterns of influenza activity seen in temperate and tropical )] TJ ET BT 35.250 734.370 Td /F1 9.8 Tf [(\(here exemplified by Colombia\) regions. The latitude of the capital city of each country is indicated in brackets in the figure )] TJ ET BT 35.250 720.634 Td /F1 9.8 Tf [(legend.)] TJ ET Q BT 26.250 682.113 Td /F1 9.8 Tf [(In the tropics, by contrast, influenza activity tends to be much more sporadic, with isolations occurring either throughout the year )] TJ ET BT 26.250 670.208 Td /F1 9.8 Tf [(or in temporally irregular outbreaks \(Fig. 1\) )] TJ ET 0.267 0.267 0.267 rg BT 213.177 671.715 Td /F4 8.7 Tf [(4)] TJ ET 0.271 0.267 0.267 rg BT 217.996 670.208 Td /F1 9.8 Tf [( . For many years the underlying reasons for influenza seasonality were widely )] TJ ET BT 26.250 658.303 Td /F1 9.8 Tf [(discussed but poorly understood )] TJ ET 0.267 0.267 0.267 rg BT 169.321 659.810 Td /F4 8.7 Tf [(5)] TJ ET BT 174.140 659.810 Td /F4 8.7 Tf [(6)] TJ ET 0.271 0.267 0.267 rg BT 178.959 658.303 Td /F1 9.8 Tf [(; recently, however, we have reported compelling evidence obtained in the guinea pig model )] TJ ET BT 26.250 646.398 Td /F1 9.8 Tf [(which suggests that seasonal variation in humidity and temperature may direct seasonal fluctuations in influenza activity )] TJ ET 0.267 0.267 0.267 rg BT 545.389 647.906 Td /F4 8.7 Tf [(7)] TJ ET BT 550.207 647.906 Td /F4 8.7 Tf [(8)] TJ ET 0.271 0.267 0.267 rg BT 555.026 646.398 Td /F1 9.8 Tf [( . In )] TJ ET BT 26.250 634.494 Td /F1 9.8 Tf [(addition, as demonstrated below, our data revealed that transmission by the aerosol route \(here defined as encompassing both )] TJ ET BT 26.250 622.589 Td /F1 9.8 Tf [(large and small droplet spread\) is acutely sensitive to both humidity and temperature, while transmission by the contact route is )] TJ ET BT 26.250 610.684 Td /F1 9.8 Tf [(not.)] TJ ET BT 26.250 591.279 Td /F4 9.8 Tf [(Sensitivity of aerosol transmission to humidity and temperature)] TJ ET BT 26.250 571.875 Td /F1 9.8 Tf [(By combining the guinea pig transmission model with the use of an environmentally controlled chamber, we have been able to )] TJ ET BT 26.250 559.970 Td /F1 9.8 Tf [(systematically evaluate the effects of temperature and relative humidity on influenza virus transmission )] TJ ET 0.267 0.267 0.267 rg BT 471.133 561.477 Td /F4 8.7 Tf [(7)] TJ ET BT 475.951 561.477 Td /F4 8.7 Tf [(8)] TJ ET 0.271 0.267 0.267 rg BT 480.770 559.970 Td /F1 9.8 Tf [( . To specifically test )] TJ ET BT 26.250 548.065 Td /F1 9.8 Tf [(the effects on transmission by the aerosol route, we used an experimental design in which infected and exposed guinea pigs )] TJ ET BT 26.250 536.160 Td /F1 9.8 Tf [(were placed in separate cages )] TJ ET 0.267 0.267 0.267 rg BT 161.190 537.668 Td /F4 8.7 Tf [(7)] TJ ET 0.271 0.267 0.267 rg BT 166.009 536.160 Td /F1 9.8 Tf [(. Each cage was open to air flow on the top and on one side and pairs of cages were placed at )] TJ ET BT 26.250 524.256 Td /F1 9.8 Tf [(a minimum distance of approximately 2 cm such that the guinea pigs were not able to touch each other. Following this )] TJ ET BT 26.250 512.351 Td /F1 9.8 Tf [(arrangement, four infected and four nave animals were placed together into an environmental chamber and monitored for virus )] TJ ET BT 26.250 500.446 Td /F1 9.8 Tf [(shedding over a period of one week. This strategy revealed a strong impact of ambient temperature and humidity conditions on )] TJ ET BT 26.250 488.541 Td /F1 9.8 Tf [(the aerosol transmission of influenza viruses \( )] TJ ET BT 226.739 488.541 Td /F4 9.8 Tf [(Fig. 2)] TJ ET BT 252.206 488.541 Td /F1 9.8 Tf [( \). In general, we found that cold and dry conditions were most favorable, )] TJ ET BT 26.250 476.637 Td /F1 9.8 Tf [(while aerosol transmission was completely blocked by warm \(30C\) or humid \(80% relative humidity\) conditions.)] TJ ET 0.965 0.965 0.965 rg 26.250 154.183 555.000 312.572 re f 0.267 0.267 0.267 rg 0.267 0.267 0.267 RG 26.250 466.756 m 581.250 466.756 l 581.250 466.006 l 26.250 466.006 l f 26.250 154.183 m 581.250 154.183 l 581.250 154.933 l 26.250 154.933 l f q 225.000 0 0 177.750 35.250 279.256 cm /I16 Do Q q 35.250 165.433 537.000 107.822 re W n 0.271 0.267 0.267 rg BT 35.250 263.732 Td /F4 9.8 Tf [(Fig. 2: Aerosol transmission of influenza A/Panama/2007/1999 virus varies with relative humidity and temperature.)] TJ ET BT 35.250 244.362 Td /F1 9.8 Tf [(Nasal wash titers of intranasally inoculated animals are indicated by dashed lines, while nasal wash titers of exposed )] TJ ET BT 35.250 230.626 Td /F1 9.8 Tf [(animals are shown with solid lines. In each experiment, n = 4 inoculated guinea pigs and 4 exposed guinea pigs. The )] TJ ET BT 35.250 216.889 Td /F1 9.8 Tf [(environmental conditions under which each experiment was performed are indicated at the top \(temperature\) and at the left )] TJ ET BT 35.250 203.153 Td /F1 9.8 Tf [(\(relative humidity, RH\). Comparison of results across each row shows the decrease in aerosol transmission with increasing )] TJ ET BT 35.250 189.417 Td /F1 9.8 Tf [(temperature; comparison of results down each column indicates the decrease in aerosol transmission with the increase in )] TJ ET BT 35.250 175.681 Td /F1 9.8 Tf [(relative humidity )] TJ ET 0.267 0.267 0.267 rg BT 108.394 178.653 Td /F4 8.7 Tf [(7)] TJ ET 0.271 0.267 0.267 rg BT 113.213 175.681 Td /F1 9.8 Tf [( .)] TJ ET Q BT 26.250 137.160 Td /F4 9.8 Tf [(Insensitivity of contact transmission to humidity and temperature)] TJ ET BT 26.250 117.755 Td /F1 9.8 Tf [(To evaluate whether transmission by the contact route would also be affected by humidity and temperature, we used an )] TJ ET BT 26.250 105.850 Td /F1 9.8 Tf [(experimental design in which infected and exposed guinea pigs were placed in the same cage together )] TJ ET 0.267 0.267 0.267 rg BT 472.810 107.357 Td /F4 8.7 Tf [(8)] TJ ET 0.271 0.267 0.267 rg BT 477.628 105.850 Td /F1 9.8 Tf [(. This set up allows for )] TJ ET BT 26.250 93.945 Td /F1 9.8 Tf [(transmission to occur by direct or indirect contact, as well as by short-range aerosol transmission. As with the aerosol )] TJ ET BT 26.250 82.041 Td /F1 9.8 Tf [(transmission experiments, environmental conditions were controlled within a chamber and exposure occurred over a period of )] TJ ET BT 26.250 70.136 Td /F1 9.8 Tf [(one week. 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In each experiment, n = 4 inoculated guinea pigs and 4 exposed guinea pigs. The )] TJ ET BT 35.250 593.229 Td /F1 9.8 Tf [(environmental conditions under which each experiment was performed are indicated above the corresponding graph )] TJ ET 0.267 0.267 0.267 rg BT 539.227 596.201 Td /F4 8.7 Tf [(8)] TJ ET 0.271 0.267 0.267 rg BT 544.046 593.229 Td /F1 9.8 Tf [(.)] TJ ET Q BT 26.250 537.510 Td /F4 12.0 Tf [(Hypothesis: differing modes of transmission predominate in temperate and tropical zones)] TJ ET BT 26.250 517.556 Td /F1 9.8 Tf [(Overall, our data in the guinea pig model indicate that aerosol transmission is acutely sensitive to relative humidity and )] TJ ET BT 26.250 505.651 Td /F1 9.8 Tf [(temperature, while contact transmission is not. An interesting hypothesis follows from these findings, namely that the )] TJ ET BT 26.250 493.746 Td /F1 9.8 Tf [(transmission of influenza viruses during the winter season in temperate regions occurs predominantly by an aerosol route, while )] TJ ET BT 26.250 481.842 Td /F1 9.8 Tf [(transmission year-round in the tropics is mainly through direct or indirect contact. If correct, this hypothesis could explain the )] TJ ET BT 26.250 469.937 Td /F1 9.8 Tf [(documented differences in seasonality between temperate and tropical regions of the world. Specifically, if viral spread in the )] TJ ET BT 26.250 458.032 Td /F1 9.8 Tf [(tropics relies mainly on fomites or direct contact, then transmission would be expected to proceed year-round despite hot and )] TJ ET BT 26.250 446.127 Td /F1 9.8 Tf [(humid climatic conditions. The persistence of influenza viruses in the human populations of the tropics is likely of great )] TJ ET BT 26.250 434.223 Td /F1 9.8 Tf [(importance to the global epidemiology of seasonal influenza: spread throughout the year in tropical regions is thought to allow )] TJ ET BT 26.250 422.318 Td /F1 9.8 Tf [(the seeding of new influenza virus strains into the Northern and Southern Hemispheres in their respective autumn months)] TJ ET 0.267 0.267 0.267 rg BT 550.274 423.825 Td /F4 8.7 Tf [(9)] TJ ET 0.271 0.267 0.267 rg BT 555.092 422.318 Td /F1 9.8 Tf [( . )] TJ ET BT 26.250 410.413 Td /F1 9.8 Tf [(Thus, the ability of influenza viruses to transmit by multiple different routes, which vary in their sensitivity to environmental )] TJ ET BT 26.250 398.508 Td /F1 9.8 Tf [(conditions, may be essential to the maintenance of these viruses in the global human population.)] TJ ET BT 26.250 379.104 Td /F4 9.8 Tf [(Summer-timer transmission of swine-origin H1N1 influenza viruses)] TJ ET BT 26.250 359.699 Td /F1 9.8 Tf [(In recent months, a very interesting exception to the standard seasonality of influenza has developed. Following its introduction )] TJ ET BT 26.250 347.794 Td /F1 9.8 Tf [(into humans in the spring of 2009, the Northern Hemisphere has seen extensive spread of a swine-origin influenza virus \(SOIV\) )] TJ ET BT 26.250 335.889 Td /F1 9.8 Tf [(over the spring and summer months )] TJ ET 0.267 0.267 0.267 rg BT 185.019 337.397 Td /F4 8.7 Tf [(10)] TJ ET 0.271 0.267 0.267 rg BT 194.656 335.889 Td /F1 9.8 Tf [(. Several possible explanations for this phenomenon exist: i\) the increased number of )] TJ ET BT 26.250 323.985 Td /F1 9.8 Tf [(summer cases is simply due to increased surveillance and testing of patients with influenza-like illness; ii\) a unique viral trait )] TJ ET BT 26.250 312.080 Td /F1 9.8 Tf [(allows SOIV strains to transmit by the aerosol route even under warm or humid conditions; iii\) unlike conventional human )] TJ ET BT 26.250 300.175 Td /F1 9.8 Tf [(strains, SOIV is able to transmit by aerosol under unfavorable conditions due to the lack of immunity against it in the human )] TJ ET BT 26.250 288.270 Td /F1 9.8 Tf [(population; or iv\) the transmission currently being seen in the Northern Hemisphere is occurring by the contact route and is )] TJ ET BT 26.250 276.366 Td /F1 9.8 Tf [(occurring with unusually high efficiency due to the low level of immunity in the population. Given that the conventional H1N1 and )] TJ ET BT 26.250 264.461 Td /F1 9.8 Tf [(H3N2 viruses which circulated in the winter of 2008/09 have not been detected out of season in high numbers, we feel that the )] TJ ET BT 26.250 252.556 Td /F1 9.8 Tf [(first explanation is most likely incorrect. The remaining hypotheses are all feasible and time and experimentation will tell which is )] TJ ET BT 26.250 240.651 Td /F1 9.8 Tf [(correct. At the present time, our favored hypothesis is the fourth, that summer-time transmission by a contact route is occurring )] TJ ET BT 26.250 228.747 Td /F1 9.8 Tf [(with increased frequency due to the antigenic novelty of the strain. If correct, we would expect increased transmission in the )] TJ ET BT 26.250 216.842 Td /F1 9.8 Tf [(coming winter months. In addition, if SOIV becomes established in the human population over the long term, then we would )] TJ ET BT 26.250 204.937 Td /F1 9.8 Tf [(expect to see subsequent epidemics fall into the conventional pattern of winter-time seasonality.)] TJ ET BT 26.250 168.335 Td /F4 12.0 Tf [(Competing interests)] TJ ET BT 26.250 148.380 Td /F1 9.8 Tf [(The authors have declared that no competing interests exist.)] TJ ET BT 26.250 111.778 Td /F4 12.0 Tf [(Acknowledgements)] TJ ET BT 26.250 91.824 Td /F1 9.8 Tf [(Figure 1 is derived from the published work of C. Viboud, W.J. Alonso and L. Simonsen.)] TJ ET BT 26.250 62.721 Td /F4 12.0 Tf [(References)] TJ ET Q q 15.000 52.291 577.500 724.709 re W n 0.965 0.965 0.965 rg 26.250 571.732 555.000 205.268 re f 0.267 0.267 0.267 rg 0.267 0.267 0.267 RG 26.250 777.000 m 581.250 777.000 l 581.250 776.250 l 26.250 776.250 l f 26.250 571.732 m 581.250 571.732 l 581.250 572.482 l 26.250 572.482 l f q 225.000 0 0 99.750 35.250 667.500 cm /I22 Do Q q 35.250 582.982 537.000 78.518 re W n 0.271 0.267 0.267 rg BT 35.250 651.976 Td /F4 9.8 Tf [(Fig. 3: Contact transmission of A/Panama/2007/1999 virus is relatively insensitive to effects of temperature and )] TJ ET BT 35.250 640.071 Td /F4 9.8 Tf [(humidity.)] TJ ET BT 35.250 620.701 Td /F1 9.8 Tf [(Nasal wash titers of intranasally inoculated animals are indicated by dashed lines, while nasal wash titers of exposed )] TJ ET BT 35.250 606.965 Td /F1 9.8 Tf [(animals are shown with solid lines. In each experiment, n = 4 inoculated guinea pigs and 4 exposed guinea pigs. The )] TJ ET BT 35.250 593.229 Td /F1 9.8 Tf [(environmental conditions under which each experiment was performed are indicated above the corresponding graph )] TJ ET 0.267 0.267 0.267 rg BT 539.227 596.201 Td /F4 8.7 Tf [(8)] TJ ET 0.271 0.267 0.267 rg BT 544.046 593.229 Td /F1 9.8 Tf [(.)] TJ ET Q BT 26.250 537.510 Td /F4 12.0 Tf [(Hypothesis: differing modes of transmission predominate in temperate and tropical zones)] TJ ET BT 26.250 517.556 Td /F1 9.8 Tf [(Overall, our data in the guinea pig model indicate that aerosol transmission is acutely sensitive to relative humidity and )] TJ ET BT 26.250 505.651 Td /F1 9.8 Tf [(temperature, while contact transmission is not. An interesting hypothesis follows from these findings, namely that the )] TJ ET BT 26.250 493.746 Td /F1 9.8 Tf [(transmission of influenza viruses during the winter season in temperate regions occurs predominantly by an aerosol route, while )] TJ ET BT 26.250 481.842 Td /F1 9.8 Tf [(transmission year-round in the tropics is mainly through direct or indirect contact. If correct, this hypothesis could explain the )] TJ ET BT 26.250 469.937 Td /F1 9.8 Tf [(documented differences in seasonality between temperate and tropical regions of the world. Specifically, if viral spread in the )] TJ ET BT 26.250 458.032 Td /F1 9.8 Tf [(tropics relies mainly on fomites or direct contact, then transmission would be expected to proceed year-round despite hot and )] TJ ET BT 26.250 446.127 Td /F1 9.8 Tf [(humid climatic conditions. The persistence of influenza viruses in the human populations of the tropics is likely of great )] TJ ET BT 26.250 434.223 Td /F1 9.8 Tf [(importance to the global epidemiology of seasonal influenza: spread throughout the year in tropical regions is thought to allow )] TJ ET BT 26.250 422.318 Td /F1 9.8 Tf [(the seeding of new influenza virus strains into the Northern and Southern Hemispheres in their respective autumn months)] TJ ET 0.267 0.267 0.267 rg BT 550.274 423.825 Td /F4 8.7 Tf [(9)] TJ ET 0.271 0.267 0.267 rg BT 555.092 422.318 Td /F1 9.8 Tf [( . )] TJ ET BT 26.250 410.413 Td /F1 9.8 Tf [(Thus, the ability of influenza viruses to transmit by multiple different routes, which vary in their sensitivity to environmental )] TJ ET BT 26.250 398.508 Td /F1 9.8 Tf [(conditions, may be essential to the maintenance of these viruses in the global human population.)] TJ ET BT 26.250 379.104 Td /F4 9.8 Tf [(Summer-timer transmission of swine-origin H1N1 influenza viruses)] TJ ET BT 26.250 359.699 Td /F1 9.8 Tf [(In recent months, a very interesting exception to the standard seasonality of influenza has developed. Following its introduction )] TJ ET BT 26.250 347.794 Td /F1 9.8 Tf [(into humans in the spring of 2009, the Northern Hemisphere has seen extensive spread of a swine-origin influenza virus \(SOIV\) )] TJ ET BT 26.250 335.889 Td /F1 9.8 Tf [(over the spring and summer months )] TJ ET 0.267 0.267 0.267 rg BT 185.019 337.397 Td /F4 8.7 Tf [(10)] TJ ET 0.271 0.267 0.267 rg BT 194.656 335.889 Td /F1 9.8 Tf [(. Several possible explanations for this phenomenon exist: i\) the increased number of )] TJ ET BT 26.250 323.985 Td /F1 9.8 Tf [(summer cases is simply due to increased surveillance and testing of patients with influenza-like illness; ii\) a unique viral trait )] TJ ET BT 26.250 312.080 Td /F1 9.8 Tf [(allows SOIV strains to transmit by the aerosol route even under warm or humid conditions; iii\) unlike conventional human )] TJ ET BT 26.250 300.175 Td /F1 9.8 Tf [(strains, SOIV is able to transmit by aerosol under unfavorable conditions due to the lack of immunity against it in the human )] TJ ET BT 26.250 288.270 Td /F1 9.8 Tf [(population; or iv\) the transmission currently being seen in the Northern Hemisphere is occurring by the contact route and is )] TJ ET BT 26.250 276.366 Td /F1 9.8 Tf [(occurring with unusually high efficiency due to the low level of immunity in the population. Given that the conventional H1N1 and )] TJ ET BT 26.250 264.461 Td /F1 9.8 Tf [(H3N2 viruses which circulated in the winter of 2008/09 have not been detected out of season in high numbers, we feel that the )] TJ ET BT 26.250 252.556 Td /F1 9.8 Tf [(first explanation is most likely incorrect. The remaining hypotheses are all feasible and time and experimentation will tell which is )] TJ ET BT 26.250 240.651 Td /F1 9.8 Tf [(correct. At the present time, our favored hypothesis is the fourth, that summer-time transmission by a contact route is occurring )] TJ ET BT 26.250 228.747 Td /F1 9.8 Tf [(with increased frequency due to the antigenic novelty of the strain. If correct, we would expect increased transmission in the )] TJ ET BT 26.250 216.842 Td /F1 9.8 Tf [(coming winter months. In addition, if SOIV becomes established in the human population over the long term, then we would )] TJ ET BT 26.250 204.937 Td /F1 9.8 Tf [(expect to see subsequent epidemics fall into the conventional pattern of winter-time seasonality.)] TJ ET BT 26.250 168.335 Td /F4 12.0 Tf [(Competing interests)] TJ ET BT 26.250 148.380 Td /F1 9.8 Tf [(The authors have declared that no competing interests exist.)] TJ ET BT 26.250 111.778 Td /F4 12.0 Tf [(Acknowledgements)] TJ ET BT 26.250 91.824 Td /F1 9.8 Tf [(Figure 1 is derived from the published work of C. Viboud, W.J. Alonso and L. Simonsen.)] TJ ET BT 26.250 62.721 Td /F4 12.0 Tf [(References)] TJ ET Q q 15.000 52.291 577.500 724.709 re W n 0.965 0.965 0.965 rg 26.250 571.732 555.000 205.268 re f 0.267 0.267 0.267 rg 0.267 0.267 0.267 RG 26.250 777.000 m 581.250 777.000 l 581.250 776.250 l 26.250 776.250 l f 26.250 571.732 m 581.250 571.732 l 581.250 572.482 l 26.250 572.482 l f q 225.000 0 0 99.750 35.250 667.500 cm /I24 Do Q q 35.250 582.982 537.000 78.518 re W n 0.271 0.267 0.267 rg BT 35.250 651.976 Td /F4 9.8 Tf [(Fig. 3: Contact transmission of A/Panama/2007/1999 virus is relatively insensitive to effects of temperature and )] TJ ET BT 35.250 640.071 Td /F4 9.8 Tf [(humidity.)] TJ ET BT 35.250 620.701 Td /F1 9.8 Tf [(Nasal wash titers of intranasally inoculated animals are indicated by dashed lines, while nasal wash titers of exposed )] TJ ET BT 35.250 606.965 Td /F1 9.8 Tf [(animals are shown with solid lines. In each experiment, n = 4 inoculated guinea pigs and 4 exposed guinea pigs. The )] TJ ET BT 35.250 593.229 Td /F1 9.8 Tf [(environmental conditions under which each experiment was performed are indicated above the corresponding graph )] TJ ET 0.267 0.267 0.267 rg BT 539.227 596.201 Td /F4 8.7 Tf [(8)] TJ ET 0.271 0.267 0.267 rg BT 544.046 593.229 Td /F1 9.8 Tf [(.)] TJ ET Q BT 26.250 537.510 Td /F4 12.0 Tf [(Hypothesis: differing modes of transmission predominate in temperate and tropical zones)] TJ ET BT 26.250 517.556 Td /F1 9.8 Tf [(Overall, our data in the guinea pig model indicate that aerosol transmission is acutely sensitive to relative humidity and )] TJ ET BT 26.250 505.651 Td /F1 9.8 Tf [(temperature, while contact transmission is not. An interesting hypothesis follows from these findings, namely that the )] TJ ET BT 26.250 493.746 Td /F1 9.8 Tf [(transmission of influenza viruses during the winter season in temperate regions occurs predominantly by an aerosol route, while )] TJ ET BT 26.250 481.842 Td /F1 9.8 Tf [(transmission year-round in the tropics is mainly through direct or indirect contact. If correct, this hypothesis could explain the )] TJ ET BT 26.250 469.937 Td /F1 9.8 Tf [(documented differences in seasonality between temperate and tropical regions of the world. Specifically, if viral spread in the )] TJ ET BT 26.250 458.032 Td /F1 9.8 Tf [(tropics relies mainly on fomites or direct contact, then transmission would be expected to proceed year-round despite hot and )] TJ ET BT 26.250 446.127 Td /F1 9.8 Tf [(humid climatic conditions. The persistence of influenza viruses in the human populations of the tropics is likely of great )] TJ ET BT 26.250 434.223 Td /F1 9.8 Tf [(importance to the global epidemiology of seasonal influenza: spread throughout the year in tropical regions is thought to allow )] TJ ET BT 26.250 422.318 Td /F1 9.8 Tf [(the seeding of new influenza virus strains into the Northern and Southern Hemispheres in their respective autumn months)] TJ ET 0.267 0.267 0.267 rg BT 550.274 423.825 Td /F4 8.7 Tf [(9)] TJ ET 0.271 0.267 0.267 rg BT 555.092 422.318 Td /F1 9.8 Tf [( . )] TJ ET BT 26.250 410.413 Td /F1 9.8 Tf [(Thus, the ability of influenza viruses to transmit by multiple different routes, which vary in their sensitivity to environmental )] TJ ET BT 26.250 398.508 Td /F1 9.8 Tf [(conditions, may be essential to the maintenance of these viruses in the global human population.)] TJ ET BT 26.250 379.104 Td /F4 9.8 Tf [(Summer-timer transmission of swine-origin H1N1 influenza viruses)] TJ ET BT 26.250 359.699 Td /F1 9.8 Tf [(In recent months, a very interesting exception to the standard seasonality of influenza has developed. Following its introduction )] TJ ET BT 26.250 347.794 Td /F1 9.8 Tf [(into humans in the spring of 2009, the Northern Hemisphere has seen extensive spread of a swine-origin influenza virus \(SOIV\) )] TJ ET BT 26.250 335.889 Td /F1 9.8 Tf [(over the spring and summer months )] TJ ET 0.267 0.267 0.267 rg BT 185.019 337.397 Td /F4 8.7 Tf [(10)] TJ ET 0.271 0.267 0.267 rg BT 194.656 335.889 Td /F1 9.8 Tf [(. Several possible explanations for this phenomenon exist: i\) the increased number of )] TJ ET BT 26.250 323.985 Td /F1 9.8 Tf [(summer cases is simply due to increased surveillance and testing of patients with influenza-like illness; ii\) a unique viral trait )] TJ ET BT 26.250 312.080 Td /F1 9.8 Tf [(allows SOIV strains to transmit by the aerosol route even under warm or humid conditions; iii\) unlike conventional human )] TJ ET BT 26.250 300.175 Td /F1 9.8 Tf [(strains, SOIV is able to transmit by aerosol under unfavorable conditions due to the lack of immunity against it in the human )] TJ ET BT 26.250 288.270 Td /F1 9.8 Tf [(population; or iv\) the transmission currently being seen in the Northern Hemisphere is occurring by the contact route and is )] TJ ET BT 26.250 276.366 Td /F1 9.8 Tf [(occurring with unusually high efficiency due to the low level of immunity in the population. Given that the conventional H1N1 and )] TJ ET BT 26.250 264.461 Td /F1 9.8 Tf [(H3N2 viruses which circulated in the winter of 2008/09 have not been detected out of season in high numbers, we feel that the )] TJ ET BT 26.250 252.556 Td /F1 9.8 Tf [(first explanation is most likely incorrect. The remaining hypotheses are all feasible and time and experimentation will tell which is )] TJ ET BT 26.250 240.651 Td /F1 9.8 Tf [(correct. At the present time, our favored hypothesis is the fourth, that summer-time transmission by a contact route is occurring )] TJ ET BT 26.250 228.747 Td /F1 9.8 Tf [(with increased frequency due to the antigenic novelty of the strain. If correct, we would expect increased transmission in the )] TJ ET BT 26.250 216.842 Td /F1 9.8 Tf [(coming winter months. In addition, if SOIV becomes established in the human population over the long term, then we would )] TJ ET BT 26.250 204.937 Td /F1 9.8 Tf [(expect to see subsequent epidemics fall into the conventional pattern of winter-time seasonality.)] TJ ET BT 26.250 168.335 Td /F4 12.0 Tf [(Competing interests)] TJ ET BT 26.250 148.380 Td /F1 9.8 Tf [(The authors have declared that no competing interests exist.)] TJ ET BT 26.250 111.778 Td /F4 12.0 Tf [(Acknowledgements)] TJ ET BT 26.250 91.824 Td /F1 9.8 Tf [(Figure 1 is derived from the published work of C. Viboud, W.J. Alonso and L. 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